长豇豆Lhc家族基因鉴定及其在盐胁迫下的表达分析

doi:10.16420/j.issn.0513-353x.2023-0786

摘要/Abstract

摘要：

捕光叶绿素a/b结合蛋白（light-harvesting chlorophyll a/b-binding protein，Lhc）直接影响光合效率和作物产量。为了系统研究长豇豆（Vigna unguiculata subsp. sesquipedalis）Lhc家族基因的特性及其在盐胁迫下的表达情况，以拟南芥Lhc蛋白序列为种子序列，并结合Lhc蛋白保守结构域PF00504，从长豇豆基因组中共鉴定了27个Lhc家族成员，并对其理化性质、基因结构、系统演化及共线性等进行分析。结果表明VuLhc启动子区光响应顺式作用元件数量远多于其他的元件，VuLhc可能参与长豇豆光调控过程，具有影响其生长发育的潜力。采用qRT-PCR检测了10个VuLhc在盐胁迫下的表达模式，结果显示其中9个呈上调趋势，其可能在豇豆对盐胁迫的响应中发挥重要作用。

关键词: 豇豆, 捕光叶绿素a/b结合蛋白, VuLhc基因家族, 表达模式, 盐胁迫

Abstract:

Light-harvesting chlorophyll a/b-binding proteins（Lhc）can directly affect photosynthetic efficiency and crop yield. To systematically study the characteristics of the Lhc gene family in long cowpea（Vigna unguiculata subsp. sesquipedalis）and their expression under salt stress，27 Lhc family members were identified from the cowpea genome using Arabidopsis Lhc protein sequences as seed sequences，combined with the conserved domain PF00504. Analyses were conducted on their physicochemical properties，gene structure，phylogenetic relationships，and synteny. The analysis indicated that the VuLhc promoter regions were enriched with light-responsive elements，significantly more than other cis-acting elements. This suggested that the VuLhc gene might be involved in the photoregulation of long cowpea，potentially influencing its growth and development. Additionally，qRT-PCR was employed to examine the expression patterns of 10 VuLhc genes under salt stress，revealing that 9 of them were upregulated，indicating their potential significant role in cowpea’s response to salt stress.

Key words: long cowpea, light-harvesting chlorophyll a/b-binding protein, VuLhc gene family, expression pattern, salt stress

王珊珊, 郭瑞, 何棱, 吴春红, 陈禅友, 万何平, 赵慧霞. 长豇豆Lhc家族基因鉴定及其在盐胁迫下的表达分析[J]. 园艺学报, 2025, 52(1): 111-122.

WANG Shanshan, GUO Rui, HE Ling, WU Chunhong, CHEN Chanyou, WAN Heping, ZHAO Huixia. Identification of Long Cowpea Lhc Gene Family and Its Expression Analysis Under Salt Stress[J]. Acta Horticulturae Sinica, 2025, 52(1): 111-122.

图/表 7

参考文献 52

[1]	Broglie R, Bellemare G, Bartlett S G, Chua N H, Cashmore A R. 1981. Cloned DNA sequences complementary to mRNAs encoding precursors to the small subunit of ribulose-1,5-bisphosphate carboxylase and a chlorophyll a/b binding polypeptide. Proceedings of the National Academy of Sciences, 78 (12):7304-7308.
[2]	Brugnoli E, Lauteri M. 1991. Effects of salinity on stomatal conductance,photosynthetic capacity,and carbon isotope discrimination of salt-tolerant (Gossypium hirsutum L.)and salt-sensitive(Phaseolus vulgaris L.)C3 non-halophytes. Plant Physiology, 95 (2):628-635. doi: 10.1104/pp.95.2.628 pmid: 16668029
[3]	Chen C J, Chen H, Zhang Y, Thomas H R, Frank M H, He Y H, Xia R. 2020. TBtools:an integrative toolkit developed for interactive analyses of big biological data. Mol Plant, 13:1194-1202.
[4]	Chen X L, Song R T, Yu M Y, Sui J M, Wang J S, Qiao L X. 2015. Cloning and functional analysis of the chitinase gene promoter in peanut. Genet Mol Res, 14 (4):12710-12722. doi: 10.4238/2015.October.19.15 pmid: 26505422
[5]	Chou K C, Shen H B. 2017. Plant-mPLoc:a top-down strategy to augment the power for predicting plant protein subcellular localization. PLoS ONE, 5:e11335.
[6]	Deng Y S, Kong F Y, Zhou B, Zhang S, Yue M M, Meng Q W. 2014. Heterology expression of the tomato LeLhcb2 gene confers elevated tolerance to chilling stress in transgenic tobacco. Plant Physiology & Biochemistry, 80:318-327.
[7]	Dong B, Wang Q Q, Zhou D, Wang Y G, Miao Y F, Zhong S W, Fang Q, Yang L Y, Xiao Z, Zhao H B. 2024. Abiotic stress treatment reveals expansin like A gene OfEXLA1 improving salt and drought tolerance of Osmanthus fragrans by responding to abscisic acid. Horticultural Plant Journal, 10 (2): 573-585.
[8]	Duvaud S, Gabella C, Lisacek F, Stockinger H, Ioannidis V, Durinx C. 2021. Expasy,the Swiss Bioinformatics Resource Portal,as designed by its users. Nucleic Acids Res, 49:W216-W227.
[9]	Feng Zhijuan, Liu Na, Zhang Guwen, Bu Yuanpeng, Wang Bin, Gong Yaming. 2024. Promoter of vegetable soybean GmDi19-3responds to salt stress and exogenous ABA and MeJA. Acta Horticulturae Sinica, 51 (12):2791-2799. (in Chinese) doi: 10.16420/j.issn.0513-353x.2023-0934
	冯志娟, 刘娜, 张古文, 卜远鹏, 王斌, 龚亚明. 2024. 菜用大豆GmDi19-3 启动子对盐胁迫和外源ABA、MeJA的响应. 园艺学报, 51 (12):2791-2799. doi: 10.16420/j.issn.0513-353x.2023-0934
[10]	Gabaldon T, Koonin E V. 2013. Functional and evolutionary implications of gene orthology. Nature Reviews Genetics, 14 (5):360-366. doi: 10.1038/nrg3456 pmid: 23552219
[11]	Ge Qianwen, Jin Baohua. 2019. Structure and function of plant light system. Botanical Research, 8 (2):170-179. (in Chinese)
	葛倩雯, 金宝花. 2019. 植物光系统的结构与功能. 植物学研究, 8 (2):170-179.
[12]	Goodstein D, Shu S, Russell H, Rochak N, Hayes R, Fazo J, Therese M, William D, Uffe H, Nicholas P, Rokhsar D. 2012. Phytozome:a comparative platform for green plant genomics. Nucleic Acids Research, 40 (D1):D1178-D1186.
[13]	Hall E A, Frate A C. 1996. Blackeye bean production in California. California:Agriculture and Natural Resources in University of California:1-23.
[14]	Han Shuang, Liu Ruixia, Zhang Zhaohe, Chen Sumei, Jiang Jiafu, Fang Weimin, Liao Yuan, Chen Fadi. 2013. Cloning of chlorophyll a/b binding protein CmLhcb1and promoter from Chrysanthemum morifolium and expression analysis. Acta Horticulturae Sinica, 40 (6):1119-1128. (in Chinese)
	韩霜, 刘瑞霞, 张兆和, 陈素梅, 蒋甲福, 房伟民, 廖园, 陈发棣. 2013. 菊花叶绿素a/b结合蛋白基因CmLhcb1及其启动子的克隆和表达分析. 园艺学报, 40 (6):1119-1128.
[15]	Jansson S. 1999. A guide to the Lhc genes and their relatives in Arabidopsis. Trends in Plant Science, 4 (6):236-240.
[16]	Jiang Q, Xu Z S, Wang F, Li M Y, Ma J, Xiong A S. 2014. Effects of abiotic stresses on the expression of Lhcb1 gene and photosynthesis of Oenanthe javanica and Apium graveolens. Biologia Plantarum, 58 (2):256-264.
[17]	Jiang Xuwen, Li Heqin, Wang Jianhua. 2015. Seed germination and physiological response of Scutellaria baicalensis seedlings to exogenous ascorbic acid under salt stress. Journal of Plant Physiology, 51 (2):166-170. (in Chinese)
	江绪文, 李贺勤, 王建华. 2015. 盐胁迫下黄芩种子萌发及幼苗对外源抗坏血酸的生理响应. 植物生理学报, 51 (2):166-170.
[18]	Kong F N, Zhou Y, Sun P P, Cao M, Li H, Mao Y X. 2016. Identification of light-harvesting chlorophyll a/b-binding protein genes of Zostera marina L. and their expression under different environmental conditions. Journal of Ocean University of China, 15:152-162.
[19]	Lescot M, Dehais P, Thijs G, Marchal K, Moreau Y, van de Peer Y, Rouze P, Rombauts S. 2002. PlantCARE,a database of plant cis-acting regulatory elements and a portal to tools for in silico analysis of promoter sequences. Nucleic Acids Research, 30:325-327.
[20]	Letunic I, Bork P. 2021. Interactive tree of life(iTOL)v5:an online tool for phylogenetic tree display and annotation. Nucleic Acids Res, 49:W293-W296.
[21]	Li Guihua, Chen Hancai, Zhang Yan. 2012. A new asparagus bean cultivar‘Fengchan 6’. Acta Horticulturae Sinica, 39 (1):197-198. (in Chinese)
	李桂花, 陈汉才, 张艳. 2012. 长豇豆新品种‘丰产6号’. 园艺学报, 39 (1):197-198.
[22]	Li X W, Zhu Y L, Chen C Y, Geng Z J, Li X Y, Ye T T, Mao X N, Du F. 2020. Cloning and characterization of two chlorophyll A/B binding protein genes and analysis of their gene family in Camellia sinensis. Scientiﬁc Reports, 10 (1):4602-4610.
[23]	Li Xianchao, Wu Yougen, Yang Dongmei, Zhang Junfeng. 2014. The response of the ultrastructure of the mesophyll cells of Hainan patchouli to salt stress. Journal of Jiangxi Agricultural University, 36 (2):300-304. (in Chinese)
	李贤超, 吴友根, 杨东梅, 张军锋. 2014. 海南广藿香叶肉细胞超微结构对盐胁迫的响应. 江西农业大学学报, 36 (2):300-304.
[24]	Liang H, Shi Q L, Li X, Gao P P, Feng D L, Zhang X M, Lu Y, Yan J S, Shen S X, Zhao J J, Ma W. 2024. Synergistic effects of carbon cycle metabolism and photosynthesis in Chinese cabbage under salt stress. Horticultural Plant Journal, 10 (2):461-472.
[25]	Liao Xiangru, He Puchao. 1996. The effect of salt stress on the H₂O₂ scavenging system in grape leaves. Acta Horticulturae Sinica, 23 (4):389-391. (in Chinese)
	廖祥儒, 贺普超. 1996. 盐胁迫对葡萄叶H₂O₂清除系统的影响. 园艺学报, 23 (4):389-391.
[26]	Liu R, Xu Y H, Jiang S C, Lu K, Lu Y F, Feng X J, Wu Z, Liang S, Yu Y T, Wang X F, Zhang D P. 2013. Light-harvesting chlorophyll a/b-binding proteins,positively involved in abscisic acid signalling,require a transcription repressor,WRKY40,to balance their function. Journal of Experimental Botany, 64 (18):5443-5456.
[27]	Livak K J, Schmittgen T D. 2001. Analysis of relative gene expression data using real-time quantitative PCR and the 2^-ΔΔCT method. Methods, 25:402-408. doi: 10.1006/meth.2001.1262 pmid: 11846609
[28]	Luo J, Abid M, Tu J, Gao P X, Wang Z P, Huang H W. 2022. Genome-wide identification of the LHC gene family in kiwifruit and regulatory role of AcLhcb3.1/3.2 for chlorophyll a content. Int J Mol Sci, 23 (12):6528-6544.
[29]	Ma Shuai, Huang Kuijian. 2023. The current situation of cowpea medication and pest control in China and suggestions for ensuring vegetable quality and safety. World Pesticides, 45 (4):13-21. (in Chinese)
	马帅, 黄魁建. 2023. 我国豇豆用药和病虫害防治现状及对保障蔬菜质量安全的建议. 世界农药, 45 (4):13-21.
[30]	Mehta P, Jajoo A, Mathur S, Bharti S. 2010. Chlorophyll a fluorescence study revealing effects of high salt stress on photosystem II in wheat leaves. Plant Physiology and Biochemistry, 48 (1):16-20. doi: 10.1016/j.plaphy.2009.10.006 pmid: 19932973
[31]	Mistry J, Chuguransky S, Williams L, Qureshi M, Salazar G, Sonnhammer E, Tosatto S, Paladin L, Raj S, Richarson L, Finn R, Bateman A. 2021. Pfam:the protein families database in 2021. Nucleic Acids Research, 49:D412-D419.
[32]	Nystrom S, McKay D. 2021. Memes:A motif analysis environment in R using tools from the MEME Suite. PLOS Comput Biol, 17:e1008991.
[33]	Shen X X, Salichos L, Rokas A. 2016. A genome-scale investigation of how sequence,function,and tree-based gene properties influence phylogenetic inference. Genome Biology and Evolution, 8:2565-2580.
[34]	Tamura K, Stecher G, Kumar S. 2021. MEGA11:molecular evolutionary genetics analysis version 11. Mol Biol Evol, 38:3022-3027.
[35]	Tang Wenli. 2008. Cloning Lhca genes from Phyllostachys pubescens and regulation of its expression at the transcriptional level by light exposure[Ph. D. Dissertation]. Beijing: China Academy of Forestry Sciences. (in Chinese)
	唐文莉. 2008. 毛竹Lhca基因的克隆和光照在转录水平对其表达的调控[博士论文]. 北京: 中国林业科学研究院.
[36]	Wang L, Wei J, Shi X Y, Qian W H, Mehmood J, Yin Y M, Jia H J. 2023. Identification of the light-harvesting chlorophyll a/b binding protein gene family in peach(Prunus persica L.)and their expression under drought stress. Genes, 14 (7):1475-1489.
[37]	Wang Y, Wang C, Rajaofera M, Zhu L, Liu W B, Zheng F C, Miao W G. 2020. WY 7 is a newly identified promoter from the rubber powdery mildew pathogen that regulates exogenous gene expression in both monocots and dicots. PLoS ONE, 15 (6):e233911.
[38]	Wang Y P, Tang H B, Debarry J, Tan X, Li J P, Wang X Y, Lee T, Jin H Z, Marler B, Guo H, Kissinger J, Paterson A. 2012. MCScanX:a toolkit for detection and evolutionary analysis of gene synteny and collinearity. Nucleic Acids Res, 40:e49.
[39]	Wang Yunhe. 2020. Identification and expression patterns of Lhc gene in ramie[M. D. Dissertation]. Wuhan: Huazhong Agricultural University. (in Chinese)
	王云鹤. 2020. 苎麻Lhc基因的鉴定和表达模式研究[硕士论文]. 武汉: 华中农业大学.
[40]	Xu Y H, Liu R, Yan L, Liu Z Q, Jiang S C, Shen Y Y, Wang X F, Zhang D P. 2012. Light-harvesting chlorophyll a/b-binding proteins are required for stomatal response to abscisic acid in Arabidopsis. Journal of Experimental Botany, 63 (3):1095-1106. doi: 10.1093/jxb/err315 pmid: 22143917
[41]	Yang Y, Wu Z K, Wu Z X, Li T Y, Shen Z, Zhou X, Wu X Y, Li G J, Zhang Y. 2023. A near-complete assembly of asparagus bean provides insights into anthocyanin accumulation in pods. Plant Biotechnology Journal, 21 (12):2473-2489. doi: 10.1111/pbi.14142 pmid: 37558431
[42]	Yu Haiying, Li Tingxuan, Zhou Jianmin. 2005. Secondary salinization of greenhouse soil and its effects on soil properties. Soils, 37 (6):581-586. (in Chinese)
	余海英, 李廷轩, 周健民. 2005. 设施土壤次生盐渍化及其对土壤性质的影响. 土壤, 37 (6):581-586.
[43]	Yuan Ruonan. 2017. Mechanism of exogenous putrescine regulation of excess excitation energy dissipation in cucumber LHC II under salt stress[Ph. D. Dissertation]. Nanjing: Nanjing Agricultural University. (in Chinese)
	袁若楠. 2017. 外源腐胺调节盐胁迫下黄瓜LHCⅡ耗散过剩激发能的机理研究[博士论文]. 南京: 南京农业大学.
[44]	Yun J F, Zhao Y, Shi F M, Wang J J. 2011. Cloning and analysis on tissue specific expression of a actin gene fragment from Agropyron mongolicum. Acta Prataculturae Sinica, 20 (2):170-176.
[45]	Zhai Yushan, Deng Yuqing, Dong Meng, Xu Qian, Cheng Guangyuan, Peng Lei, Lin Yanquan, Xu Jingsheng. 2016. Cloning and expression of ScLhca3,a light harvesting chlorophyll a/b binding protein gene in sugarcane. Journal of Crops, 42 (9):1332-1341. (in Chinese)
	翟玉山, 邓宇晴, 董萌, 徐倩, 程光远, 彭磊, 林彦铨, 徐景升. 2016. 甘蔗捕光叶绿素a/b结合蛋白基因ScLhca3的克隆及表达. 作物学报, 42 (9):1332-1341.
[46]	Zhang Haoying, Yang Yating, Zhang Zhaoyuan, Jiang Peng, Zhu Shidong, Wang Chenggang, Hou Jinfeng, Chen Guohu, Tang Xiaoyan, Wang Wenjie, Wu Jianqiang, Yuan Lingyun. 2023. Identification and expression analysis of the Lhca gene family in Chinese cabbage. Molecular Plant Breeding, 15:1-11. (in Chinese)
	张豪英, 杨雅婷, 张钊源, 江鹏, 朱世东, 汪承刚, 侯金锋, 陈国户, 唐小燕, 王文杰, 吴建强, 袁凌云. 2023. 白菜Lhca基因家族的鉴定与表达分析. 分子植物育种, 15:1-11.
[47]	Zhang Q, Ma C, Wang X, Ma Q, Fan S, Zhang C. 2021. Genome-wide identiﬁcation of the light-harvesting chlorophyll a/b binding(Lhc)family in Gossypium hirsutum reveals the influence of GhLhcb2.3 on chlorophyll a synthesis. Plant Biology(Stuttgart,Germany), 23 (5):831-842.
[48]	Zhang Y, Raza A, Huang H, Su W, Luo D, Zeng L, Ding X Y, Cheng Y, Liu Z F, Li Q A, Lv Y, Zou X L. 2022. Analysis of Lhcb gene family in rapeseed(Brassica napus L.)identifies a novel member“BnLhcb3.4”modulating cold tolerance. Environ Exp Bot, 198:104848.
[49]	Zhao Qi, Ru Jingna, Li Yitong, Wang Chao, Xu Zhaoshi, Wang Ruihui. 2022. Identification and expression pattern analysis of wheat Lhc gene family. Journal of Plant Genetic Resources, 23 (6):1766-1781. (in Chinese) doi: 10.13430/j.cnki.jpgr.20220401001
	赵奇, 茹京娜, 李宜统, 王超, 徐兆师, 王睿辉. 2022. 小麦Lhc基因家族鉴定与表达模式分析. 植物遗传资源学报, 23 (6):1766-1781.
[50]	Zhao S, Gao H B, Luo J W, Wang H B, Dong Q L, Wang Y P, Yang K Y, Mao k, Ma F W. 2020a. Genome-wide analysis of the light-harvesting chlorophyll a/b-binding gene family in apple(Malus domestica) and functional characterization of Md Lhcb4.3,which confers tolerance to drought and osmotic stress. Plant Physiology and Biochemistry, 154:517-529.
[51]	Zhao Y G, Kong H, Guo Y L, Zou Z. 2020b. Light-harvesting chlorophyll a/b-binding protein-coding genes in jatropha and the comparison with castor,cassava and Arabidopsis. Peer J, 8 (4):e8465.
[52]	Zou Z, Yang J H. 2019. Genomics analysis of the light-harvesting chlorophyll a/b-binding(Lhc) superfamily in cassava(Manihot esculenta Crantz). Gene, 702 (51):171-181.

基因名称 Gene name	正向引物（5′-3′） Forward primer	反向引物（5′-3′） Reverse primer
VuLhc1	CGGAAAGGTAGAGCTGGTGG	CTCTGGGTCAAGTTCTGCGT
VuLhc3	TTCGACCCTCTGGGACTAGG	CCAGCCTCAAACCATGCAAC
VuLhc12	CCCTGCCTCCTTCAAGACTG	GCAAGCTCATCATTGGCAGG
VuLhc14	GCAAACTCGGTGTTGCAGTT	GGTAGCGATCCATCAAGCCA
VuLhc15	TGTTTGCGATGCTCGGTTTC	TCTCAGCTGATCCAGCAAGC
VuLhc16	TCACCAAAGGTGAGCACGAA	CTGCCCTTTGGCTACTTCCA
VuLhc22	CCGGTAAACTGAACCGGGAA	GCCACTCTCCTTTCTTGGCT
VuLhc23	CCAAGGTTGTAAAGCCAAAGCA	CCCAACAATGATGCCGCAAA
VuLhc24	AACAGCGACCTACATGGCTC	GGAATGAACCAGCTCTGCCT
VuLhc25	AAGCCTTGTCGGTGACTACG	TTCTTCGCCAGGTTCTGGTC
Actin	AATGATCGGAATGGAAGCTG	TGGAATGTGCTGAGAGATGC

基因名称 Gene name	正向引物（5′-3′） Forward primer	反向引物（5′-3′） Reverse primer
VuLhc1	CGGAAAGGTAGAGCTGGTGG	CTCTGGGTCAAGTTCTGCGT
VuLhc3	TTCGACCCTCTGGGACTAGG	CCAGCCTCAAACCATGCAAC
VuLhc12	CCCTGCCTCCTTCAAGACTG	GCAAGCTCATCATTGGCAGG
VuLhc14	GCAAACTCGGTGTTGCAGTT	GGTAGCGATCCATCAAGCCA
VuLhc15	TGTTTGCGATGCTCGGTTTC	TCTCAGCTGATCCAGCAAGC
VuLhc16	TCACCAAAGGTGAGCACGAA	CTGCCCTTTGGCTACTTCCA
VuLhc22	CCGGTAAACTGAACCGGGAA	GCCACTCTCCTTTCTTGGCT
VuLhc23	CCAAGGTTGTAAAGCCAAAGCA	CCCAACAATGATGCCGCAAA
VuLhc24	AACAGCGACCTACATGGCTC	GGAATGAACCAGCTCTGCCT
VuLhc25	AAGCCTTGTCGGTGACTACG	TTCTTCGCCAGGTTCTGGTC
Actin	AATGATCGGAATGGAAGCTG	TGGAATGTGCTGAGAGATGC

基因ID Gene ID	基因名称 Gene name	氨基酸数 No. of amino acids	相对分子质量/kD Mw	等电点 pI	平均疏水性 Gravy	亚细胞定位 Subcellular localization
Vigun01g226400.1	VuLhc1	279	30.70	6.01	-0.022	叶绿体Chloroplast
Vigun02g173500.1	VuLhc2	264	28.07	5.13	-0.020	叶绿体Chloroplast
Vigun03g133000.1	VuLhc3	258	27.48	6.15	0.142	叶绿体Chloroplast
Vigun03g145400.1	VuLhc4	263	27.91	5.29	-0.007	叶绿体Chloroplast
Vigun04g103500.2	VuLhc5	307	33.85	7.20	-0.066	叶绿体Chloroplast
Vigun04g128900.1	VuLhc6	264	28.00	5.15	-0.003	叶绿体Chloroplast
Vigun04g129000.1	VuLhc7	264	28.01	5.15	-0.013	叶绿体Chloroplast
Vigun04g129200.1	VuLhc8	173	18.34	4.79	0.212	叶绿体Chloroplast
Vigun04g129300.1	VuLhc9	264	28.06	5.05	-0.011	叶绿体Chloroplast
Vigun04g129500.1	VuLhc10	264	28.01	5.15	-0.020	叶绿体Chloroplast
Vigun04g131200.1	VuLhc11	264	28.09	5.29	-0.014	叶绿体Chloroplast
Vigun04g167600.1	VuLhc12	289	30.80	5.72	0.022	叶绿体Chloroplast
Vigun05g198900.1	VuLhc13	269	28.88	5.25	-0.016	叶绿体Chloroplast
Vigun06g224500.1	VuLhc14	258	27.36	6.75	0.139	叶绿体Chloroplast
Vigun07g168400.1	VuLhc15	288	30.93	6.35	0.018	叶绿体Chloroplast
Vigun07g234800.1	VuLhc16	183	19.60	9.39	0.178	叶绿体Chloroplast
Vigun08g000700.1	VuLhc17	342	37.88	6.66	-0.016	细胞膜/线粒体/过氧化物酶体 Cell membrane/mitochondria/ peroxisome
Vigun08g216300.1	VuLhc18	265	28.67	5.46	-0.131	叶绿体Chloroplast
Vigun08g221400.1	VuLhc19	276	29.72	7.87	-0.074	叶绿体Chloroplast
Vigun09g003500.1	VuLhc20	270	28.91	6.42	-0.004	叶绿体Chloroplast
Vigun09g040000.1	VuLhc21	261	28.95	5.04	-0.075	细胞膜Cell membrane
Vigun09g075200.1	VuLhc22	252	27.85	6.97	-0.137	叶绿体Chloroplast
Vigun09g165900.1	VuLhc23	269	28.51	6.63	0.252	叶绿体Chloroplast
Vigun09g238500.1	VuLhc24	264	28.58	8.63	0.055	细胞膜/线粒体 Cell membrane/ mitochondria
Vigun10g078500.1	VuLhc25	289	31.26	5.79	-0.120	叶绿体Chloroplast
Vigun10g155200.1	VuLhc26	269	28.87	6.08	0.052	叶绿体Chloroplast
Vigun11g049900.1	VuLhc27	269	29.03	5.24	0.041	叶绿体Chloroplast

基因ID Gene ID	基因名称 Gene name	氨基酸数 No. of amino acids	相对分子质量/kD Mw	等电点 pI	平均疏水性 Gravy	亚细胞定位 Subcellular localization
Vigun01g226400.1	VuLhc1	279	30.70	6.01	-0.022	叶绿体Chloroplast
Vigun02g173500.1	VuLhc2	264	28.07	5.13	-0.020	叶绿体Chloroplast
Vigun03g133000.1	VuLhc3	258	27.48	6.15	0.142	叶绿体Chloroplast
Vigun03g145400.1	VuLhc4	263	27.91	5.29	-0.007	叶绿体Chloroplast
Vigun04g103500.2	VuLhc5	307	33.85	7.20	-0.066	叶绿体Chloroplast
Vigun04g128900.1	VuLhc6	264	28.00	5.15	-0.003	叶绿体Chloroplast
Vigun04g129000.1	VuLhc7	264	28.01	5.15	-0.013	叶绿体Chloroplast
Vigun04g129200.1	VuLhc8	173	18.34	4.79	0.212	叶绿体Chloroplast
Vigun04g129300.1	VuLhc9	264	28.06	5.05	-0.011	叶绿体Chloroplast
Vigun04g129500.1	VuLhc10	264	28.01	5.15	-0.020	叶绿体Chloroplast
Vigun04g131200.1	VuLhc11	264	28.09	5.29	-0.014	叶绿体Chloroplast
Vigun04g167600.1	VuLhc12	289	30.80	5.72	0.022	叶绿体Chloroplast
Vigun05g198900.1	VuLhc13	269	28.88	5.25	-0.016	叶绿体Chloroplast
Vigun06g224500.1	VuLhc14	258	27.36	6.75	0.139	叶绿体Chloroplast
Vigun07g168400.1	VuLhc15	288	30.93	6.35	0.018	叶绿体Chloroplast
Vigun07g234800.1	VuLhc16	183	19.60	9.39	0.178	叶绿体Chloroplast
Vigun08g000700.1	VuLhc17	342	37.88	6.66	-0.016	细胞膜/线粒体/过氧化物酶体 Cell membrane/mitochondria/ peroxisome
Vigun08g216300.1	VuLhc18	265	28.67	5.46	-0.131	叶绿体Chloroplast
Vigun08g221400.1	VuLhc19	276	29.72	7.87	-0.074	叶绿体Chloroplast
Vigun09g003500.1	VuLhc20	270	28.91	6.42	-0.004	叶绿体Chloroplast
Vigun09g040000.1	VuLhc21	261	28.95	5.04	-0.075	细胞膜Cell membrane
Vigun09g075200.1	VuLhc22	252	27.85	6.97	-0.137	叶绿体Chloroplast
Vigun09g165900.1	VuLhc23	269	28.51	6.63	0.252	叶绿体Chloroplast
Vigun09g238500.1	VuLhc24	264	28.58	8.63	0.055	细胞膜/线粒体 Cell membrane/ mitochondria
Vigun10g078500.1	VuLhc25	289	31.26	5.79	-0.120	叶绿体Chloroplast
Vigun10g155200.1	VuLhc26	269	28.87	6.08	0.052	叶绿体Chloroplast
Vigun11g049900.1	VuLhc27	269	29.03	5.24	0.041	叶绿体Chloroplast